Ramalina pusiola Müller Argoviensis

Marcano, Vicente, Méndez, Antonio Morales & Prü, Ernesto Palacios, 2021, The genus Ramalina Acharius (Ascomycota, Lecanoromycetes, Ramalinaceae) in northern South America, Phytotaxa 504 (1), pp. 1-77 : 54-57

5. Ramalina pusiola Müller Argoviensis

Flora, Jena 73: 338 (1890) . Type:― TANZANIA. Süden von Usumbara: Tumakauya, Dr Hans Meyer, sept. 1888 (lectotype-Stein 12 WRSL, isotype G).

Thallus fistulose, corticolous, rigid, ascendent, pale yellow, irregularly and densely branched, up to 5.0 cm long, growing from a broad holdfast, producing crystals on the upper surface (Fig. 22). Branches dorsiventral, hollow, sometime partly compressed, unevenly inflated, often having perforations at the tips, surface smooth. Pseudocyphellae not seen. Soralia not seen. Chondroid tissue not cracked, continuous. Pycnidia not seen.Apothecia rounded, subterminal to terminal, concave, spurred, up to 4 mm in diameter, pale or reddish brown. Ascospores uniseptate, ellipsoid or fusiform, 15–18 µm x 4.5–5.5 µm.

Chemistry (TLC,HPTLC):This species presents8strains which are summarized in Table8.Strain1. Homosekikaic, norstictic, ramalinolic (tr.), 4ˊ– O –demethylsekikaic and sekikaic acids. Strain 2. Homosekikaic and sekikaic acids (Morales 155–89). Strain 3. Homosekikaic, sekikaic and boninic acids (López-Figueiras 17152–B; Marcano 6109, Morales 122–89, 143–89). Strain 4. Homosekikaic, sekikaic, boninic, fumarprotocetraric, succinprotocetraric and protocetraric acids (Morales 135–89). Strain 5. Homosekikaic, sekikaic, 4ˊ– O –demethylsekikaic, boninic, fumarprotocetraric, succinprotocetraric and protocetraric acids (Marcano 148–92, Marcano 149–92, Morales 147– 89). Strain 6. Homosekikaic, sekikaic, ramalinolic and boninic acids (Cleef & van der Hammen 4961). Strain 7. Homosekikaic, sekikaic, 4ˊ– O –demethylsekikaic and boninic acids (Vareschi 4499, Aguirre & Sipman 5592). Strain 8. Homosekikaic, sekikaic, ramalinolic and 4ˊ– O –demethylsekikaic acids (Vareschi 6200, 9841).

Ecology and distribution: Ramalina pusiola is common and widespread in northern South America where it ranges from lowland to alpine altitudes. It is most abundant in lower montane forest and secondary forest at 1200 to 3800 m where it is epiphytic, on tree trunks and branches.Associated lichens in the montane forest include Teloschistes flavicans , Usnea durietzii Motyka (1938: 503), Ramalina cumanensis and R. celastri . In Colombia it is found growing on Ficus sp. and fruit trees in coffee plantations. It is known from East Africa ( Ethiopia, Kenya, Tanzania, Uganda) and northern South America ( Brazil, Colombia and Venezuela).

Remarks: Ramalina pusiola is morphologically similar to R. calcarata but differs in having well-defined, orbicular to elongate, perforations on the lower surface; non spurred apothecia; long ellipsoid to fusiform spores; and sekikaic acid and its aggregates in the medulla. By contrast, Ramalina calcarata exhibits fibrose-reticulate perforations; spurred apothecia; ellipsoid spores (12–15 x 5–7 µm) and divaricatic, salazinic and protocetraric acids in the medulla. Ramalina pusiola produces homosekikaic, sekikaic and boninic, acids as major substances; and ramalinolic, 4ˊ– O – demethylsekikaic acids and the protocetraric acid complex as accessory substances (Table 8). In East Africa ( Krog & Swinscow 1974) R. pusiola synthesize sekikacic acid and aggregates while Kashiwadani & Kalb (1993) reported the presence of homosekikaic, sekikaic, 4ˊ– O –methylnorhomosekikaic and norstictic acids (+/–) in the 15 collections of this species they examined.

Specimens examined: COLOMBIA: Risaralda: Municipio Santa Rosa de Cabal, vereda El Limbo, Finca Los Lajos , 1600 m, 20 September 1984, J . Aguirre & H . Sipman 5592 ( B) . Cundinamarca: Sasaima, finca San José , 1850 m, 20 July 1972, A . Cleef & Th. van der Hammen 4961 ( B) . VENEZUELA: Aragua: Near Colonia Tovar, 2000 m, Vareschi 9841 ( VEN) . Miranda: El Volcán, 1400 m, 5 January 1957, Vareschi 6200 ( VEN); El Volcán, 1430 m, 1 March 1956, Vareschi 4499 ( VEN) . Mérida: Los Topes, Chiguará , 1200 m, 15 April 1988, A . Morales 122, 135, 143, 147 ( MER); Las Playitas, near Bailadores , 1950 m, 14 April 1992, V . Marcano 3–92, 6101 ( MER); San Juanito, Chiguará , 1200 m, 15 March 1989, A . Morales 155–89 ( MER); Santo Domingo, Paramo de Mucubají , 3500–3800 m, March 1992, V . Marcano 148–92, 149–92 ( MER) . Táchira: Near Pregonero, Garcia Lake , 1900 m, 14 April 1992, V . Marcano 6106, 6107, 6109, 6110 ( MER); near Paramo Portachuelo, 2800–2900 m, 14 April 1992, V . Marcano 6111 ( MER); Betania, Tama valley, near Villa Paez , 2400–2500 m, 14 April 1992, V . Marcano 6113 ( MER); near Villa Paez, Delicias , 1900–2100 m, 13 April 1992, V . Marcano 6116– A ( MER) .

. Ramalina rectangularis Nylander (Fig. 37)

Bull. Soc. Linn. Normandie, ser. 2, 4: 121 (1870) . Type:― BRAZIL. Bahia, Blanchet (lectotype H–NYL 37509, syntypes PC, UPS, W).

Thallus corticolous, pendulous and dichotomously branched, pale yellow, up to 15 cm long, branching dichotomic anisotomic. Branches solid, rectangular in cross-section, rigid, flattened, not contorted, 2–3 mm wide, surface smooth, showing protuberances marginal. Isidial structures verruciformis, disposed along the margin of branches. Pseudocyphellae marginal, linear, concave or convex, continuous. Cortex distinct, 14–21 μm thick. Chondroid tissue distinctly cracked, continuous. Medulla whitish, loose. Pycnidia and apothecia not seen.

Chemistry (TLC, HPTLC): Strain 1. Usnic acid only (Vareschi 5504; Sipman & Escobar 33849, Vareschi 5405). Strain 2. Salazinic acid (Morales 94, 95; Vareschi 3388, 5448). Strain 3. Norstictic, stictic (tr.) and salazinic acids (Marcano et al. 7217).

Ecology and distribution: This species is found growing on shrubs and branches, associated with Stereocaulon Hoffman (1796: 128) , Hypotrachyna ( Vainio 1890: 38) Hale (1974: 340) [ Everniastrum Hale ex Sipman (1986: 237) ] and crustose lichens, in cloud forests and subparamo at 1250–3350 m. In Colombia it was found growing on rocks in a pastures on a steep slope. It is only known from South America ( Colombia, Brazil, Venezuela).

Remarks: Ramalina rectangularis is a rare South American species, previously known only from the type collection from State of Bahiate, Northeastern Brazil ( Kashiwadani & Kalb 1993, Gumboski 2016). It is characterized by a pendulous and dichotomously branched thallus, a smooth surface showing marginal protuberances, and marginal, verruciformis isidial structures. Ramalina rectangularis resembles R. puiggari , but the latter species has a canaliculate and more flexible thallus, and lacks medullary substances. Ramalina rectangularis could also be confused with R. usnea , but this species has a contorted thallus (in the study area) and contains sekikaic acid and its aggregates. Ramalina rectangularis exhibits three chemical strains, one which lacks medullary substances and two with the βorcinol depsidones, norstictic, stictic and salazinic acids. Ramalina rectangularis from Brazil was reported to contain salazinic acid only ( Kashiwadani & Kalb 1993, Gumboski 2016).

Specimens examined: COLOMBIA: Antioquia: Municipio Medellín , El Boquerón, mountain pass, c. 10 km NW of Medellín on road to Santa Fe de Antioquia, 2500 m, 26 June 1986, H . Sipman & M . Escobar 33849 ( B) . VENEZUELA: Mérida: La Culata, Mucumpís road, 2800–3200 m, October 1993, V . Marcano , R . Vidal & A . Morales 7217 ( MER); El Paramito, Chiguara, 1250 m, 12 February 1984, A . Morales 94, 95 ( MERF; B) . Miranda: Los Guayabitos, near la Mata , 1300 m, 20 June 1954, V . Vareschi 3388 ( VEN) . Táchira: Paramo El Batallón, 3100–3350 m, 3 October 1956, Vareschi 5504, 5405 ( VEN); Paramo El Pulpito, 3140–3300 m, 3 October 1956, V . Vareschi 5448 ( VEN) .

. Ramalina reducta Krog & Swinscow var. reducta

Norw. J. Bot. 23: 172 (1976). Type :― ETHIOPIA. Province Bale : Dinshu, Tapper 923 (holotype BM, isotype O).

Thallus corticolous, erect, dichotomously branched, 2–4 cm high. Branches compact, flattened, 2–3,5 cm wide, palmate, surface rugose, without a lower cortex (Fig. 16). Pseudocyphellae tuberculate, ellipsoid, sorediate, arising on the upper surface, 400–500 μm long, 300–350 μm wide. Soralia forming coarse granules. Chondroid tissue distinctly cracked. Medulla dense, exposed on the lower side where cartilaginous tissue forms a reticular structure. Pycnidia and apothecia not seen.

Chemistry (TLC, HPTLC): Strain 1. 2ˊ– O –demethylpsoromic and psoromic acids (Cleef 4849–C, 9984–E; Lopez-Figueiras &. Morales 29648; Marcano 6630, 6635, 6636, 6637). Strain 2. Psoromic acid (Sipman 23612; Sipman & Hollaender 34497).

Ecology and distribution: This species is an epiphyte in exposed habitats, growing on the branches of trees and shrubs where it is associated with R. celastri , R. pusiola , R. dictyota , Usnea spp., Hypotrachyna caraccensis ( Taylor 1847: 163) Hale (1975: 26) , H. ducalis ( Jatta 1908: 407) Hale (1975: 33) , Oropogon (Theodor Magnus Fries 1861: 49) spp. and Teloschistes flavicans . In Colombia R. reducta var. reducta grows on shrubs and branches in very wet Weinmannia , Vallea stipularis Linnaeous and Miconia forests well as branches of Espeletia at 1000–3885 m (Fig. 30). It is known from Australia, East Africa ( Ethiopia, Tanzania, Uganda, Transvaal) and South America ( Colombia and Venezuela).

Remarks: The morphological and chemical characters of this species in Venezuela have been described in detail by Marcano & Morales (1993b). Ramalina reducta has two varieties, one characterized by the presence of psoromic acid ( var. reducta ) and the other characterized by the presence of norstictic acid (sometimes with psoromic; var. colombiana ; see below) as major medullary substances. Both varieties have disjunct distributions and occur in separate populations. In northern South America Ramalina reducta var. reducta is known mostly from Venezuela whereas var. colombiana is known only from Colombia. In East Africa ( Krog & Swinscow 1976) and Australia ( Stevens 1987) this species produces psoromic acid only. Ramalina reducta resembles R. tucumanensis Räsänen (1949: 46) , but the latter species lacks soralia and pseudocyphellae, has a different chemistry (viz. sekikaic acid as major medullary substance), and it is known only from Perú and Argentina ( Kashiwadani 1987).

Specimens examined: COLOMBIA: Bogota: Paramo de Chisaca , along road Usme-Nazareth, 3500 m, 14 July 1986, H . Sipman & M . Hollaender 34497 ( B) . Cundinamarca: San Francisco, vereda Sabaneta, near Cueva Grande, 2500 m, 17 July 1986, H . Sipman, H . Cardozo & M . Ballestreros 23612 ( B); Valle Teusacá , 5 km SW La Calera, 2850 m, 9 July 1972, A . Cleef & M . Cleef – van Rens 4849–c ( B) . Boyaca: Sierra Nevada del Cocuy , Alto de la Cueva, 3885 m, 5 June 1973, A. M . Cleef 9984–e ( B). Valle: Cali, University of Valle , 1000 m, 1986, J . Rubiano s/n– A ( B) . VENEZUELA: Mérida: Paramo La Negra , 3000 m, M . Lopez-Figueiras & Dana Griffin III 31582 ( MERF); Paramo Las Lajas , 3200–3300 m, M . Lopez-Figueiras et A . Morales 29648 ( MERF); Monte Zerpa, Serranía La Culata , 2200–2400 m, V . Marcano 5557 ( MERF); Las Playitas, near Bailadores , 1950 m, V . Marcano 6630, 6635, 6636, 6637 ( MERF) . Táchira: Laguna García, near Pregonero, 1900 m, V . Marcano 6601 ( MERF) . Trujillo: Paramo El Morro, 2300 m, M . Lopez-Figueiras 28231 ( MERF) .

. Ramalina reducta var. colombiana Marcano & Morales var. nov.

Thallus ut in Ramalina reducta var. reducta sed acidum norsticticum continente differt.

Type:― COLOMBIA. Cundinamarca: near Valley of Neusa river, 3 km Southern, 26 May 1972, A . Cleef & R . Jaramillo 4190–b (holotype B) .

Chemistry ( TLC, HPTLC): Strain 1. Psoromic, norstictic and hypoprotocetraric acids (Cleef & R. Jaramillo 4190–b). Strain 2. Norstictic and salazinic acids (Sipman & Valencia 10297– A). Strain 3. Norstictic and connorstictic acids (Sipman & Reyes 34453).

Ecology and distribution: This variety is found growing on branches and trunks of Xylosma in forests with Weinmannia and Vallea at 2700–2950 m. It is known only from Colombia.

Remarks: Ramalina reducta var. colombiana is distinguished of var. reducta by the presence of norstictic acid as the major medullary substance. This varieties shows three chemically distinct strains, all with various β-orcinol depsidones as accessory substances.

Additional specimens examined: COLOMBIA. Boyaca: Municipio Cucaita, ca. 10 km W of Tunja, alomng road to Villa de Leiva , 2950 m, 12 July 1986, H . Sipman & O . Reyes 34453 ( B) . Cundinamarca: Between Bogotá and Fusagasuga, c 5 km SW of roadtoll, 2700 m, 31 January 1979, H . Sipman & H . Valencia 10297– A ( B, COL, U) .

. Ramalina rigida Persoon ex Acharius

Syn. Lich.: 294 (1814). Type:―ST. DOMINGO (holotype H – ACH 1822 ) .

Thallus corticolous, decumbent. Branches terete or subterete, solid, 0.8–0.25 µm broad; surface smooth; apices tapering, often ending in granular soredia. Pseudocyphellae marginal, laminal, ellipsoid, punctiform, usually becoming sorediate. Soralia terminal or laminal. Cortex distinct, chondroid tissue cracked. Pycnidia not seen. Apothecia rare, subterminal, lateral, showing pseudocyphellate margins, epithecium 4–5 µm thick, hymenium 36–40 µm thick. Ascospores 1–septate, curved, 10–11 x 3–3.5 µm.

Chemistry (TLC, HPTLC): Strain 1. Homosekikaic (tr.), sekikaic (tr.) and salazinic acids (Vareschi 3448). Strain 2. Stictic and salazinic acids (Vareschi 6179).

Ecology and distribution: This species is found growing on Cereus near shore, on Leguminosae in dry forests, at 50–1400 m. It is known from the Caribbean (type collection) and South America ( Brazil and Venezuela). Krempelhuber (1876c) recorded this species as corticolous from collections carried out by Glaziou (no. 1860, 1861, 1863–1865) in the Brazilian Amazonas, near Rio Janeiro, while Kashiwadani & Kalb (1993) reported the species as saxicolous from collections in Rio Janeiro by Vainio distributed in his Lichenes Brasilienses Exsiccati.

Remarks: Ramalina rigida is rare in northern South America. It could be confused with R. tenella , R. dendriscoides and R. sorediosa , but these species have corticolous, erect and mostly shrubby thalli. Ramalina santanensis is also similar but has a knotty surface and lacks soralia. Ramalina rigida shows two distinct chemical strains with salazinic acid as the major medullary substance. In Brazil and the Caribbean ( Kashiwadani & Kalb 1993) this species contains salazinic acid only.

Specimens examined: VENEZUELA: Miranda: Los Guayabitos , 1200–1400 m, 1 January 1957, V . Vareschi 6179 ( VEN) . Falcón: Near Tucacas , 50–100 m, 4 July 1954, V . Vareschi 3448 ( VEN) .